Reproductive condition and ovarian atresia in the common kilka (Clupeonella cultriventris) from the Caspian Sea years after a dramatic outbreak of the comb jelly fish (Mnemiopsis leidyi)


  • Narjes Karimi Fisheries Department, Faculty of Natural Resources, University of Guilan
  • Ali Bani Department of Biology, Faculty of Science, University of Guilan,
  • Shahram Abdolmalaki International Sturgeon Research Institute



Caspian Sea, Clupeonella cultriventris, atresia, group-synchronous, spawning season, somatic condition, Mnemiopsis leidyi


To assess the reproductive condition of the common kilka, Clupeonella cultriventris, the somatic condition and reproductive characteristics were studied by histological analysis. A total of 800 specimens from the Bandar Anzali coast on the Caspian Sea were sampled. The gonadosomatic index (GSI) for females and males attained maximum values in May in the first period of sampling and in March-April in the second period, suggesting temporal variations in spawning activity. The pattern of oocyte frequency distribution demonstrated that the common kilka is a batch spawner with group-synchronous oocyte development. The estimated relative batch fecundity was 6718 oocytes g–1. A low level of atresia (4.7%) during the spawning season showed that this species was in optimal somatic condition. This weakens the hypothesis that abundance of the ctenophore (Mnemiopsis leidyi) in the Caspian Sea, as a food rival for kilka, influences the body condition and subsequently the reproductive capability of this species.


Download data is not yet available.


Asahina K., Hanyu I. 1983. Role of temperature and photoperiod in annual reproductive cycle of the rose bitterling Rhodeus ocellatus ocellatus. Nippon. Suisan. Gakk. 49: 61-67.

Balon E. K. 1984. Life histories of Arctic Charrs: an epigenetic explanation of their invading ability and evolution. In: Johnson L., Burns B.L., (eds), Biology of the Arctic Charr, Proceedings of the International Symposium on Arctic Charr, University of Manitoba Press, Winnipeg, pp. 109-141.

Besharat K., Khatib S. 1993. Determinate of the commercial catch region in the Iranian coastal zone, 1990-1991. Final report, Mazandaran Fisheries Research Center. 150 pp. (In Persian).

Bilio M., Niermann U. 2004. Is the comb jelly really to blame for it all? Mnemiopsis leidyi and the ecological concerns about the Caspian Sea. Mar. Ecol. Pog. Ser. 269: 173-183.

Brown-Peterson N.J., Overstreet R.M., Lotz J.M., Franks J.S., Burns K.M. 2001. Reproductive biology of cobia, Rachycentron canadum, from coastal waters of the southern United States. Fish. Bull. 99: 15-28.

Brown-Peterson N.J., Wyanski D.M., Saborido-Rey F., Macewicz B.J., Lowerre-Barbieri S.K. 2011. A standardized terminology for describing reproductive development in fishes. Mar. Coast. Fish. 3: 52-70.

Conover D.O. 1990. The relation between capacity for growth and length of growing season: evidence for and implications of countergradient variation. Trans. Am. Fish Soc. 119: 416-430.<0416:TRBCFG>2.3.CO;2

Davis T.L.O., West G.J. 1993. Maturation, reproductive seasonality, fecundity, and spawning frequency in Lutjanus vittus (Quoy and Gaimard) from the North West Shelf of Australia. Fish. Bull. 91: 224-236.

Einum S., Fleming I.A., Cote I.M., Reynolds J.D. 2003. Population stability in salmon: species effect of population size and female reproductive allocation. J. Anim. Ecol. 72: 811-821.

Fazli H., Besharat K. 1998. Kilka fishes stock assessment with fishing zones monitoring and hidroacostic method. Fisheries Research Center of Mazandaran Province, 150 pp. (In Persian)

Fazli H., Borani M.S., Janbaz A.A. 2000. The biological characteristics of Clupeonella cultriventris caspia in southern coast and the impacts of Mnemiopsis leidyi invasion on the Caspian ecosystem. Pajohesh. Sazandgi. 69: 87-96. (In Persian)

Fazli H. 2007. Population dynamics and stock assessment of kilka (genus: Clupeonella) in Iranian waters of the Caspian Sea. PhD thesis. Pukyong National University, South Korea. 145 pp.

Fowler A.J., Mcleay L., Short D.A. 2000. Spatial variation in size and age structures and reproductive characteristics of the King Gorge whiting (Percoidi: Sillahinidae) in South Australian waters. Aust. J. Mar. Freshw. Res. 51: 423-429.

Ganias K., Somarakis S., Machias A., Theodorou A. 2004. Pattern of oocyte development and batch fecundity in the Mediterranean sardine. Fish Res. 67:13-23.

Hayes J.P., Shonkwiler J.S. 2001. Morphometric indicators of body condition: worthwhile or wishful thinking? In: Speakman J.R. (ed.), Body composition analysis of animals. Cambridge University Press, Cambridge, pp. 8-38. PMid:11348409

Hunter J.R., Macewicz B.J. 1985a. Measurement of spawning frequency in multiple spawning fishes. NOAA. Technical. Report. NMFS. 36: 79-94.

Hunter J.R., Macewicz B.J. 1985b. Rates of atresia in the ovary of captive and wild northern anchovy, Engraulis mordax. Fish. Bull. 83: 119-136.

Iranian Fisheries Organization. 2011. Year book of Iranian Fisheries Organization statistics for 1994-2010. Research and Development sector of Iranian Fisheries Organization. Tehran, 65 pp. (In Persian).

Iran Metrological Organization. [cited May 2009]. Available from URL:

Jakob E.M., Marshall S.D., Uetz G.W. 1996. Estimating fitness: A comparison of body condition indices. Oikos 77: 61-67.

Janbaz A.A. 2006. Population dynamics and stock assessment of common kilka (Clupeonella cultriventris) in Iranian waters of the Caspian Sea. MSc thesis. Islamic Azad University Lahijan, Iran, 125 pp. (In Persian)

Kideys A.E., Ghasemi S., Ganinejad D., Roohi A., Bagheri S. 2001. Strategy for combating Menemiopsis in the Caspian waters of Iran. Final Report, July 2001, Caspian Environment Programme, Baku, Azerbaijan.

Kjesbu O.S. 1994. Time of start of spawning in Atlantic cod (Gadus morhua) females in relation to vitellogenic oocyte diameter, temperature, fish length and condition. J. Fish. Biol. 45: 719-735.

Kjesbu O.S., Solemdal P., Bratland P., Fonn M. 1996. Variation in annual egg production in individual captive Atlantic cod (Gadus morhua). Can. J. Fish. Aquat. Sci. 53: 610-620.

Kruse G.H., Tyler A.J. 1983. Simulation of temperature and upwelling effects on the English sole (Parophrys vetulus) spawning season. Can. J. Fish. Aquat. Sci. 40: 230-237.

Lam T.J. 1983. Environmental influences on gonadal activity in fish. Fish. Physiol. 9B: 65-115.

Lambert Y., Dutil J.D. 2000. Energetic variability in prediction by cod, Cadus morhua in relation to spawning level of somatic energy reserves. Can. J. Fish. Aquat. Sci. 57: 815-825.

Lowerre-Barbieri S. K., Brown-Peterson N. J., Murua H., Tomkiewicz J., Wyanski D., Saborido-Rey F. 2011a. Emerging issues and methodological advances in fisheries reproductive biology. Mar. Coast. Fish. 3: 32-51.

Lowerre-Barbieri S.K., Chittenden Jr. M.E., Barbieri L.R. 1996. The multiple spawning pattern of weakfish in the Chesapeake Bay and Middle Atlantic Bight. J. Fish. Biol. 48: 1139-1163.

Lowerre-Barbieri S.K., Ganias K., Saborido-Rey F., Murua H., Hunter J.R. 2011b. Reproductive timing in marine fishes: variability, temporal scales, and methods. Mar. Coast. Fish. 3: 71-91.

Marshall C.T., Kjesbu O.S., Yaragina N.A., Solemdal P., Ulltang Ø. 1998. Is spawner biomass a sensitive measure of the reproductive and recruitment potential of Northeast Arctic cod? Can. J. Fish. Aquat. Sci. 55: 1766-1783.

Miranda A.C.L., Bazzoli N., Rizzo E., Sato Y. 1999. Ovarian follicular atresia in two teleost species: a histological and ultrastructural study. Tissue. Cell. 31: 480-488. PMid:18627868

Murua H., Saborfido-Rey F. 2003. Female reproductive strategies of marine fish species of the north atlantic. J. Northw. Atl. Fish. Sci. 33: 23-31.

Naderi M., Fazli H., Afraee M., Ganjian A. 1997. The study of reproduction, fecundity and diet in 3 species of kilka in the southern part of the Caspian Sea (Babolsar region). Iran J. Fish Sci. 6: 65-74.

Pianka E.R. 2000. Evolutionary ecology. Sixth edition. Benjamin-Cummings, Addison-Longman, San Francisco, 528 pp.

Roohi A., Zulfigar Y., Kideys A., Aileen T., Eker-Develi E., Ganjian A. 2008. Impact of a new invasive ctenophore (Mnemiopsis leidyi) on the zooplankton community of the Southern Caspian sea. Mar. Ecol.-Evol. Persp. 29: 421-434.

Pourgholam R., Sedov V., Yermalchev V., Besharat K., Fazli H. 1996. Stock assessment of kilka fishes by hydro acoustic method, 1994-1995. Final report, Mazandaran Fisheries Research Center, 125 pp. (In Persian)

Prikhodko B.I. 1981. Ecological features of the Caspian kilka (Genus Clupeonella). Scripta Publishing Co. pp. 27-35.

Rideout R.M., Rose G.A., Burton M.P.M. 2005. Skipped spawning in female iteroparous fishes. Fish Fish. 6: 50-72.

Rideout R.M., Tomkiewicz J. 2011. Skipped spawning in fishes: more common than you might think. Mar. Coast. Fish. 3: 176-189.

Rijnsdorp A. D. 1991. Changes in fecundity of female North Sea plaice (Pleuronectes platessa L.) between three periods since 1900. ICES J. Mar. Sci. 48: 253-280.

Rizzo E., Bazzoli N. 1995. Follicular atresia in curimata-pioa Prochilodus affinis Reinhardt, 1874 (Pisces, Characiformes). Rev. Bras. Biol. 55: 697-703.

Roff D.A. 1992. The evolution of life history, theory and analysis. Chapman and Hall, New York, 535 pp.

Scott S.G., Pankhurst N.W. 1992. Interannual variation in the reproductive cycle of the new zealand snapper Pagrus auratus (Sparidae). J. Fish. Biol. 41: 685-696.

Trippel E.A., Kjesbu O.S., Solemdal P. 1997. Effects of adult age and size structure on reproductive output in marine fishes. In: Chabmers R.C., Trippel E.A. (eds), Early life history and recruitment in fish populations, pp. 31-62.

Tyler C.R., Sumpter J.P. 1996. Oocyte growth and development in teleosts. Rev. Fish. Biol. Fisher. 6: 287-318.

Wallace R.A., Selman K., Greeley M.J., Begovac P.C., Lin Y.W., Mcpherson R., Petrino T.R. 1987. Current status oocyte growth. In: Idler D.R., Crim L.W., Walsh J.M. (eds), International symposium on reproductive physiology of fish. St John's: Memorial University of Newfoundland, pp. 167-177.

Ware D.M. 1984. Fitness of different reproductive strategies in teleosts fishes. In: Potts G.W., Wootton R.J. (eds), Fish reproduction: strategies and tactics. Academic Press, New York, pp. 349-366.

Wootton R.J. 1982. Environmental factors in fish reproduction. In: Richter C.J.J., Goss H.J.T. (eds), Reproductive physiology of fish. Pudoc, Wageningen, NL, pp. 210-219.

Wootton R.J. 1984. Introduction: tactics and strategies in fish reproduction. In: Pots G.W., Wootton R.J. (eds), Fish reproduction: strategies and tactics. Academic Press, New York, pp. 1-12.

Yoneda M., Tokimura M., Fujita H., Takeshita N., Takeshita K., Matsuyama M., Matsuura S. 2001. Reproductive cycle, fecundity, and seasonal distribution of the anglerfish Lophius litulon in the East China and Yellow seas. Fish. Bull. 99: 356-370.




How to Cite

Karimi N, Bani A, Abdolmalaki S. Reproductive condition and ovarian atresia in the common kilka (Clupeonella cultriventris) from the Caspian Sea years after a dramatic outbreak of the comb jelly fish (Mnemiopsis leidyi). scimar [Internet]. 2013Sep.30 [cited 2023Dec.11];77(3):501-9. Available from: